Collective Decision-making In Honey Bees: How Colonies .
Behav Ecol Sociobiol(1991) 28:277-290Behavioral Ecologyand Sociobiology Springer-Verlag1991Collective decision-making in honey bees:how colonles choose among nectar sourcesThomas D. Seeley 1, Scott Camazine 1, and James Sneyd 2.t Section of Neurobiologyand Behavior, Mudd Hall, CornellUniversity,Ithaca, NY 14853, USA2 Centre for MathematicalBiology,MathematicalInstitute, 24-29 St. Giles', Oxford OX1 3LB,UKReceived April 3, 1990 / Accepted September 30, 1990Summary. A honey bee colony can skillfully chooseamong nectar sources. It will selectively exploit the mostprofitable source in an array and will rapidly shift itsforaging efforts following changes in the array. Howdoes this colony-level ability emerge from the behaviorof individual bees ? The answer lies in understanding howbees modulate their colony's rates of recruitment andabandonment for nectar sources in accordance with theprofitability of each source. A forager modulates its behavior in relation to nectar source profitability: as profitability increases, the tempo of foraging increases, theintensity of dancing increases, and the probability ofabandoning the source decreases. How does a foragerassess the profitability of its nectar source? Bees accomplish this without making comparisons among nectar sources. Neither do the foragers compare differentnectar sources to determine the relative profitability ofany one source, nor do the food storers compare different nectar loads and indicate the relative profitabilityof each load to the foragers. Instead, each forager knowsonly about its particular nectar source and independently calculates the absolute profitability of its source. Eventhough each of a colony's foragers operates with extremely limited information about the colony's foodsources, together they will generate a coherent colonylevel response to different food sources in which betterones are heavily exploited and poorer ones are abandoned. This is shown by a computer simulation of nectar-source selection by a colony in which foragers behaveas described above. Nectar-source selection by honeybee colonies is a process of natural selection among alternative nectar sources as foragers from more profitablesources "survive" (continue visiting their source) longerand ' reproduce" (recruit other foragers) better than doforagers from less profitable sources. Hence this colonialdecision-making is based on decentralized control. We* P r e s e n t address." Department of Biomathematics,UCLA Schoolof Medicine, 10833 La Conte Avenue, Los Angeles, CA 90024,USAO f f p r i n t requests to : T.D. Seeleysuggest that honey bee colonies possess decentralizeddecisi0n-making because it combines effectiveness withsimplicity of communication and computation within acolony.IntroductionRecently, numerous authors have noted that coloniesof advanced social insects constitute higher-order cognitive entities, that is, supraorganismal systems capableof evaluating situations and producing adaptive solutions to problems (Hofstadter 1979; Markl 1985; Seeleyand Levien 1987; Wilson and H611dobler 1988; Franks1989). Examples of this colony-level problem-solving include a swarm of honey bees selecting the best nest sitefrom a dozen or more possibilities (Lindauer 1955; Seeley 1982), a colony of honeypot ants assessing thestrength of a neighboring colony and deciding accordingly whether or not to attack (H611dobler 1981 ; Lumsden and H611dobler 1983), and a colony of army antschoosing a bearing for each day's swarm raid so as toavoid the previous day's foraging zone (Franks andFletcher 1983; Deneubourg et al. 1989). All such ensemble cognitive performances present us with the puzzleof how the problem-solving abilities of a colony arisefrom the actions and interactions of the insects composing the colony.We address this mystery in the context of honey beecolonies choosing among patches of flowers, selectivelyexploiting the most profitable ones. The ecology of thisdecision-making can be summarized as follows (reviewedin Seeley 1985). Each year a colony of bees consumesconsiderable food, approximately 20 kg of pollen and60 kg of honey. The thousands of foragers in a colonylaboriously gather this food from the flower patches dotting the countryside around the colony's hive. Typicallya colony will know each day about a dozen or morepotential food sources, each with its own level of profit-
278ability, determined by such variables as the distancefrom the hive and the abundance and quality of thefood. To gather its food efficiently, a colony must deployits foragers among the flower patches in accordance withtheir profitabilities.We begin by documenting the skill with which a colony chooses among nectar sources so that most of itsnectar comes always from the richest sources. Then weshow how this selective exploitation arises through precise modulation of the rates of recruitment and abandonment for different nectar sources in relation to their profitabilities. This points out the crucial question: howare the rates of these two processes adaptively regulated ?After describing the multidimensional modulation offoraging behavior in relation to nectar source profitability, we experimentally analyze how each forager assessesnectar source profitability. The two principal competinghypotheses are that each forager judges by itself the profitability of its nectar source, or that each forager isgiven this information by the food storer bees in thehive. Next we check our understanding of how the jointactions of a colony's foragers result in producing colonylevel decision-making by formulating a mathematicalmodel of this process and comparing the decision-making patterns of simulated and real colonies. We end bydiscussing the general implications of our investigation.applied to its abdomen. Finally, the bees were gently poured intoa cage containing a sugar water feeder and the mother queen.This procedure was repeated several times a day for 2 days until4000 bees were labelled.At the end of the second day, the labelled workers and theirqueen were transferred to a two-frame observation hive identicalto the one described in Seeley (1989). The bottom frame containeda mixture of brood (eggs and larvae), honey, and pollen, and thetop frame was empty except for a few cells of pollen. On 4 Junethe colony was moved to Cranberry Lake and left undisturbedfor 4 more days, though bees were trained to the feeders to setup the experimental array and to provide food for the colony.On the evening of 8 June, the colony's population was censusedby counting the number of bees inside 10 randomly selected 2.5 x2.5 cm grid squares on each side of the observation hive: the colonycontained approximately 3450 bees. Rainy weather prevailed forthe next 10 days, during which all we could do was provide foodat the feeders whenever the bees could fly. This kept the colonyfed and ensured that the dozen or so bees trained to each feederwould continue visiting the feeders. By the time the weather cleared(19 June) the colony had grown noticeably; fortunately the foragerswere still all labelled bees. A census on the evening of 19 Juneindicated a population of approximately 4200 bees.MethodsMeasuring forager group size, and recruitment and abandonmentrates. We recorded which bees visited each food source during1. Colony-level analysis: selective exploitation of nectarsourcesevery half-hour period throughout two 8-h sets of observations.We accomplished this using roll call sheets listing the identificationcodes of all 4000 bees and crossing off the identification codesof all bees seen at a feeder during each half hour. Each sheettherefore showed which individuals visited a particular feeder atleast once during a 30-rain period, and how many in total didso. This number we call the "forager group size." Comparisonsbetween roll call sheets for a feeder also revealed when (to thenearest 30 rain) individuals were first sighted at (i.e., were recentlyrecruited to) the feeder or were last sighted at (i.e., were soonto abandon) the feeder. From these data we calculated the percapita recruitment rate or abandonment rate for each feeder foreach half hour by dividing the number of recruits or deserterscounted during that half hour by the forager group size for theprevious half hour.A colony composed of individually identifiable bees was presentedwith two nectar sources with different profitabilities. The colony'sdifferential exploitation of the two sources was measured by determining the total number of different individuals visiting each foodsource. Also, the different rates of recruitment and abandonmentfor the two feeders, which ultimately generated the exploitationdifference between the two food sources, were determined fromrecords of each bee's visits to the feeders.Study site. All observations were made at the Cranberry Lake Biological Station (44009 ' N, 74 48 ' W) in northern New York State.This site is surrounded by forests, bogs, and lakes for more than20 km in all directions, hence forage for bees is extremely sparseand no native honey bees survive there. A full-size colony, kepton platform scales, lost weight (0.50 0.30 kg) on 15 days andgained weight (0.15 kg) on just 1 day during the 16-day studyperiod in June 1989. The scarcity of food enabled us to controlthe forage collected by our study colony even though its foragersflew freely from the hive.Study colony. We assembled a colony of individually identifiablebees over a 2-day period, 1-2 June 1989, in Ithaca, as follows.First, at the beginning of each day, approximately 2500 workerbees from a colony of Italian honey bees (Apis mellifera ligustica)were shaken into a wire cage. From this, we periodically shookgroups of approximately 50 bees into plastic bags and placed themin a refrigerator to immobilize them. After at least 15 min of cooling, the bees in a bag were removed from the refrigerator andpoured onto a container of "blue ice", where they stayed chilledduring the labelling operation. A bee tag (Opalithpl/ittchen; with500 number and color combinations) was glued on the thorax ofeach bee, and a dot of one of eight different colors of paint wasFeeders and experimental layout. The feeders were pneumatic feeding dishes identical to those described in a previous paper (Seeley1989). The observation hive and the two feeders were placed inan approximately linear arrangement, with the hive in the centerand the feeders 400 m to the north and south, in sinai1 clearings.Both food sources contained the scent anise in a vented reservoirbeneath the feeder and also in the sucrose solution (60 gl of aniseextract per liter).2. Individual-level analysis: behavior modulationin relation to nectar source profitabilityTwo colonies were established in adjacent observation hives and30 bees from each colony were trained to visit separate feedersat the same distance from the hives. The profitability of one (theexperinaental) colony's feeder was systematical(y changed duringthe day and the behavioral adjustments of its foragers were recorded. MeanwNle, the profitability of the other (the control) colony's feeder was left unchanged and the behaviors of its foragerswere recorded as a check for possible confounding changes in theambient conditions.Study site, bees, experimental layout, and feeders. This experimentwas conducted in July 1987 in connection with other experimentsalready published (Seeley 1989). It involved the same study site,the same two colonies in observation hives, the same experimentallayout, and the same feeders as were described in the prior report.
279The only difference between the present experiment and the onereported in Fig. 2 of Seeley 1989 was that, in the present case,the fullness of the experimental colony's honey combs was keptconstant and the profitability of it's feeder was varied, whereas,before, the fullness of the experimental colony's honey combs wasvaried and the profitability of it's feeder was kept constant.Measuring recruitment and recording the behaviors of foragers. Recruitment from each colony was measured by training 30 beesto a feeder using standard techniques (von Frisch 1967), labelingthese bees (the recruiters) with individually identifiable paintmarks, and counting the number of unlabeled bees (the recruits)appearing at the feeder every 15 rain. Each recruit was capturedin a plastic bag (to minimize release of alarm pheromone) shortlyafter its arrival at the feeder, and was frozen at the end of theday.Roll calls of the labeled bees at 30-min intervals indicated thenumber of recruiters at each feeder; this number dropped below30 for the experimental colony when the profitability of its feederwas lowered and some bees abandoned it. The probability of visiting the feeder was calculated by dividing the number of labeledbees visiting the feeder by 30, that is, the number of labeled beesthat visited the feeder when all did so.Measurements were also made of several variables of the inhive behavior of the labelled bees from the experimental colony.This involved following one (randomly selected) labelled bee ata time from when it entered the hive to when it exited it. Onevariable was the "time to start of unloading," which is the timeinterval between entering the hive and beginning to transfer nectarto a food storer bee. The "time to end of unloading" is the timeinterval between entering the hive and completing the transfer ofnectar to food storers. The "time in hive after unloading" is thetime interval between finishing unloading nectar and leaving thehive. Another variable was "dance circuits per bee per return tothe hive," which was measured by counting the dance circuits eachbee performed during its time inside the hive. The "probabilityof dancing" was calculated by dividing the number of bees thatdanced by the total number of bees that were followed (i.e., samplesize). Finall , the "foraging tempo," measured in units of tripsto the feeder per bee per 30 min, was calculated by first dividingthe number of labelled bees visiting the feeder by the number oflabelled bees entering the hive per rain (measured with ten l-raincounts of labelled bees coming in the entrance). This quotient indicates the average time per roundtrip to the feeder. Next, 30 minwas divided by the roundtrip time to determine the number oftrips to the feeder that each bee was making in 30 rain.Statistical test. A single classification ANOVA (Sokal and Rohlf1981) was performed for each variable for which there was anestimate of the mean and variance, i.e., all variables except forthe probability of visiting the feeder, the probability of dancing,and foraging tempo.3. Individual-level analysis: assessing nectar sourceprofitabilityTwo feeders were established, one with a relatively dilute sugarsolution but located near the hive, and the other with a moreconcentrated sugar solution but positioned far from the hive. Beeswere trained to forage from each feeder, and their behaviors insidethe hive, especially the strength of their recruitment dances, weremonitored to determine which feeder was judged more profitable.The answer reveals whether it is the food storers or the foragersthat assess nectar source profitability.Study site, bees, experimental layout, and feeders. This experimentwas performed in Ithaca, New York, using a small colony of Italianhoney bees in a two-frame observation hive housed inside a laboratory building. The hive and feeders were identical to those describedin a previous study (Seeley 1989). Both feeders were positionedalong a rarely used road (one 50 m and the other 1250 m fromthe hive) that traverses flat, treeless agricultural fields; hence, bothfeeders had virtually identical exposure to wind and sun. Fifteenbees were trained to each feeder using standard techniques (vonFrisch 1967). Each bee was labeled with paint marks coding individual identity and which feeder she was visiting. Recruits to eachfeeder were captured in plastic bags to prevent forager build-upat the feeders.Recording the behaviors of foragers. Labelled foragers were followed one at a time inside the observation hive from time of arrivalto departure. Measurements were made, as previously described,of the time to start of unloading, the dance circuits per bee perreturn to the hive, and the probability of dancing.Statistical tests. We used t-tests, either Student's or the test forequality of two proportions using arcsine transformations (Sokaland Rohlf 1981).Results1. Colony-level analysisPattern: selective exploitation o f nectar sources. F i g u r e 1shows t h a t w h e n a c o l o n y was given a series o f choicesb e t w e e n t w o feeders w i t h different profitabilities, it c o n sistently f o c u s e d its c o l l e c t i o n efforts o n the m o r e p r o f i t able feeder. T h e net result was t h a t the c o l o n y s t e a d i l yt r a c k e d the richest source o f n e c t a r in the c h a n g i n g array. N o t e t h a t the m e a s u r e o f f o r a g i n g effort s h o w nin Fig. 1 - the t o t a l n u m b e r o f different i n d i v i d u a l s visiting the feeder in a h a l f h o u r - u n d e r e s t i m a t e s the colo n y ' s r e s p o n s e differential to the two feeders. This isbecause the p o o r e r feeder (0.75 mol/1), relative to thericher feeder (2.50 tool/l), was n o t o n l y visited b y fewerbees, b u t also was visited at a l o w e r rate b y each bee(see d a t a on f o r a g i n g t e m p o in Table 1).Process: tuning the rates o f recruitment and abandonment. Because the size o f a g r o u p is d e t e r m i n e d b y theh i s t o r y o f a d d i t i o n s to a n d s u b t r a c t i o n s f r o m the g r o u p ,it was clear a p r i o r i t h a t the c h a n g e s in f o r a g e r g r o u psize s h o w n in Fig. 1 reflect c h a n g e s in the rates o f recruitment (additions) and abandonment (subtractions) foreach feeder. T h e c o m p l e t e p i c t u r e o f the r e c r u i t m e n t a n da b a n d o n m e n t d y n a m i c s for the two feeders is s h o w nin Fig. 2. O n the m o r n i n g o f 19 June, there was essentially no r e c r u i t m e n t to o r a b a n d o n m e n t o f the n o r t h feeder(1.00 mol/1), a n d the f o r a g e r g r o u p size here rose o n l yto 12 bees, n a m e l y the ones t h a t h a d f o r a g e d here thep r e v i o u s day. A t the s o u t h feeder (2.50 mol/1), h o w e v e r ,there was s t r o n g r e c r u i t m e n t , especially e a r l y in them o r n i n g w h e n the feeder was still lightly e x p l o i t e d , a n dlittle a b a n d o n m e n t , so the f o r a g e r g r o u p size i n c r e a s e dd r a m a t i c a l l y , rising f r o m 0 to 91 bees in 4 h o u r s . T h e nin the a f t e r n o o n , the s o u t h feeder ( n o w 0.75 tool/l) received negligible r e c r u i t m e n t b u t intense a b a n d o n m e n t ,so t h a t its f o r a g e r g r o u p size fell b a c k to j u s t 10 beesb y the e n d o f the a f t e r n o o n . M e a n w h i l e , the n o r t h feeder( n o w 2.50 tool/l) h a d b e c o m e the t a r g e t o f s t r o n g recruitm e n t b u t little a b a n d o n m e n t , with the net result t h a t
280North FeederHiveSouth Feeder TT(9 8 AM1.00 M2.50 M.: ::.LUZ- TNoon2.50 M 0.75 M :o.'. . ::.o o O o eO o o o oOo eeee ooe@ oeOOo oeoeoO ::::: ooeo oooooe00000 eel looooe 0 o oo eeo o TO O l4 PMGT 8 AMT f (92. Individual-level analysisT(90.75 M2.50 M.LUz .- NoonT(3 T 0.75 M4 PM2.50 Myt3unexpectedly large forager group size - as m a n y as 32bees - for the poorer (south, 0.75 mol/1) feeder on themorning of 20 June. This morning surge in foragers tothe poorer feeder arose because bees that had foragedhere the previous morning came out to reconnoitre thesite of their prior success, not because bees were recruiting nestmates to the 0.75 re l/1 solution at this feeder.Virtually all of the bees (46 out of 48) that visited thesouth feeder on the morning of 20 Jun
Thomas D. Seeley 1, Scott Camazine 1, and James Sneyd 2. t Section of Neurobiology and Behavior, Mudd Hall, Cornell University, Ithaca, NY 14853, USA 2 Centre for Mathematical Biology, Mathematical Institute, 24-29 St. Giles', Oxford OX1 3LB, UK Received April 3, 1990 / Accepted September 3
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